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Gastropoda: Architectonicidae ) ' The Feeding, Larval D ispersal, and Metamorphosis of Philippia (Gastropoda: Architectonicidae ) ' ROB ERT ROB ERTSON, RUDOLF S. SCHELTEMA, and FRANK W . ADAMS2 ABSTRACT: In the Hawaiian Islands, Philippia (Psilaxis) radiata ( Reding) lives in sand or rubble near the hermatypic stony coral Porites lobata Dana, and emerges to feed at night on the polyps, Other species of Philippia (Psilaxis) probably have the same mode of life with corals. Philippia (Psilaxis ) veliger lar­ vae are abundant in tropical and subtropical oceanic plankton distant from any potential shallow-water hosts, and are dispersed great distances by near-surface currents. Duration of the pelagic larval stage is between several weeks and 6 months or longer. Metamorphosis, involving loss of the 4-10bed velum, initial growth of the teleoconch, and other changes, can precede contact with a host and is induced by capture from the plankton. Newly settled Philippia quickly attain a stage of arrested growth and can remain alive without feeding for several months . At this stage the postla rvae presumably crawl in search of hosts, and failure to find hosts doubtless causes the high mortality observed. Experiments at Woods H ole, Massachusetts, with newly metamorphosed P. (Psilaxis) krebsii (March), obtained as larvae from plankton in the Sargasso Sea, together with the ahermatypic coral Astrangia danae Agassiz, reveal physical prob­ lems for Philippia in assuming the adult mode of life with other corals. Young Philippia showed no ability to detect A strangia except by touch. Young Philippia lacked immuni ty to Astrangia nematocysts but were not seriously injured by them. The young gastropods are, however, subject to predation by this coral. Most con­ tacts with the living tissues of Astrangia caused a Philippia to be promptly drawn through the mouth and ultimately digested. The large protoconchs of Psilaxis would preclude their being swallowed by hermatypic corals such as Porites, with polyps smaller than those of A strangia. Proboscis eversion and feeding were not observed in young P. krebsii. IN A PAPER reporting that the genus H eliacus ters. However, Nordsieck has informed us (in comprises obligate symbionts that feed on co­ litt.) that he has no more specific data than lonial zoanthiniarian sea anemones, Robertson that P. hybl'ida lives in a coral and coralline (1967) mentioned that nothing was known of habitat. Taylor (1968, p. 187) mentioned that the feeding habits of the two other major gen­ Arcbitectonica perspectioa (Linn.) "probably era in the Architectonicidae, Philippia and Ar­ feeds upon zoanthids" at 15 to 25 meters in cbitectonica. Subsequently, Nordsieck (196 8, the Seychelles, but Taylor has informed us (in p. 65) has mentioned the Mediterranean and lit!.) that he dredged A. perspectiua together Lusitanian species Philippia (Philippia) hy­ with zoanthids, and extrapolated from Robert­ brida (Linn.) "auf Koral len" at about 40 me- son's (1967) conclusions on Heliacus, Shuto (1969, p. 25, footnote) states that A. perspec­ 1 Woods Hole Oceanographic "Institution Contri ­ tiva is "strictly speaking . synbiotic [sic] bution No . 2312. Manuscript received May 9, 1969. with sand-dwelling sea anemones" ; no sup­ 2 Addr esses, respectively: Academy of N atural Sci­ porting data were given. Abbott (1968 , p. 86) ences of Phi ladelphia, Philadelphia, Pennsylvania mentioned Arcbitectonica nobilis Reding' "in 19103; Woods H ole Oceanographic Institution, Woods Hole, Massachusetts 02543; and 1739C Ala shallow water among sea pansies." Arcbitec­ Moana Boulevard, Honolulu, Hawaii 96815. tonica nobilis commonly is washed ashore on 55 56 PACIFIC SCIENCE, Vol. 24, January 1970 ," .. ,.. ... ,t) '~" ....-.... , i "2; \ >tt~ ~ ~~ ., ~'t:" ':"';'~ > : '.::~>:,.\~?:~.~ :. ' ,: . 5 mm . FIG. 1. Postlarval Philippia (Psilaxis) radiata (Roding ) feeding on the polyps of the hermatypic stony coral Porites lobata Dana. Compos ite drawing by Mary Fuges from photographs of a Hawaiian animal feeding, from other photographs of extended Philippia animals, and from specimens of the shell and coral. The Pbilipp i« lives in the coarse sand near the coral and crawls out to feed at night. Philippia (Feeding, etc.) ---':"'ROBERTSON ET AL. 57 Texas beaches together with pennatulids Our information on larval dispersal and the (Renilla muelleri Kolliker; teste R. T. Abbott) . metamorphosis of Philippia enhance the in­ Most recently, A. nobilis has several times been terest of the nutritional dependence of the post­ seen with its proboscis everted, presumably larvae on coelenterates. All three species in the while feeding on something (Futch, 1969). subgenus Psilaxis have planktotrophic veligers Thus, all five of these tantalizing reports pro­ that remain for long periods in the plankton vide no definite information. There are simi­ (Robertson, 1964 ; Scheltema, 1968; Robert­ larly vague hints that Gyriscas, a fourth archi­ son, 1970b) . We have but scanty data on the tectonicid genus, may live with gorgonians two Indo-Pacific species, and therefore we sum­ (Tiberi, 1868) and sponges (Powell, 1965, marize here some of our data on the larvae of pp. 161-162). the Atlantic Ocean species Philippia (Psilaxis) New observations (by Adams) are here re­ erebsii (March) , on its metamorphosis in the ported on Philippia (Psilaxis) radiata (Ro­ laboratory at Woods Hole, and on the inter­ ding) which in the Hawaiian Islands lives action between the young postlarvae and the near and feeds on the soft tissues of the herma­ ahermatypic coral Astrangia danae Agassiz. typic stony coral Porites lobata Dana (Fig. 1). Information on the systematics, zoogeography, (This discovery was briefly mentioned in Ha­ and bathymetry of Indo-Pacific Philippia (Psi­ waiian Shell News, vol. 15, no. 10, p. 4; Oc­ laxis) , and on eggs and young postlarvae in tober 1967) . The Architectonicidae become the umbilicus of adults, is given in the follow­ the fifth family of prosobranch gastropods ing paper (Robertson, 1970b) . definitely known to include predators or para­ sites on stony (scleractinian) corals (Robert­ PROBOSCIS STRUCTURE son, 1970a). The habitat and feeding data are preceded here by a review of the structure of Like other architectonicids, Philippia has a the proboscis, jaws, and radula of Philippia. remarkably long, acrembolic proboscis. When Their specialization and uniformity lead us to this is fully everted, the true mouth and the believe that food and mode of feeding are uni­ small buccal mass containing the radula are at form throughout the genus. the tip. Thus the animal can feed only when FIGS . 2 and 3. Shell of the PhiZippia radiat a specimen seen to have fed on Porites Zobata (45 feet off Makua, western Oahu, Hawaiian Islands ; shell in the collection of Frank W . Adams) . Apical and basal views, respect ively (both X 6) . 58 PACIFIC SCIENCE, Vol. 24, January 1970 the proboscis is fully everted. The lining of the genetic relationship and by Thiele (1928, pp . buccal cavity and esophagus is cuticularized as 87- 88) to convergence. Epitoniids are now in epitonii ds, and this prevents injury from known to feed on coelenterates exclusively and nematocysts in both groups (Robertson, 1970b). we are inclined to agree with Thiele. The laterally paired jaws of Philippia are elon­ gate, are comprised of scalelike elements, and Philipp ia radiata: are the distal part of the buccal cavity wall. HAWAIIAN HABITAT AND F EEDING DATA Upon complete proboscis eversion, the jaws are near the mouth edge. The radula, uniform Philippia (Psilaxis) radiata is one of two throughout the genus, is narrow and comprised 'closely related species in the H awaiian Islands of elonga te teeth bent distally and with one to (Robertson, 1970b) . The postlarvae, rarely six points. As observed long ago by T roschel collected alive, are known from shallow water (1875, p. 155), these architectonicid radu lar to about 150 feet. During daylight, the ani­ teeth resemble those of epitoniids. The resem­ mals are usually buried 1 to 4 inches in silty blance was attributed by Troschel to phylo- sand or loose, algal-covered rubble, the sand FIG. 4. Porites /obata. Some of the calices of the cleaned skeleton of the coral seen to have been fed on by Pbilippi« radiate. X 30. Philippia (Feeding, etc.) - ROBERTSON ET AL. 59 and rubbl e shallowly overlying basalt (Ha­ boscis fully everted adjacent to the Porites col­ waiian Shell N ews, vol. 16, no. 12, p. 7; ony as it fed delicately on the still partly December 1968) . In every case known to us, expanded polyps. Under bright photoflood the living animals were near massive or incrust­ lights, two color photographs were taken be­ ing colonies of Porites lobata. fore the Philipp ia inverted its proboscis and One living Philippia radlata with a fairly retracted into its shell. Th e drawing (F ig. 1), small shell ( Figs. 2 and 3) was collected by based in part on these photographs, probably Adams while scuba diving at a depth of 45 does not show the full length of the extended feet off Makua, western Oahu, in April or proboscis. May 1967. It was taken from a patch of sand near a small living colony of Porites lobata on SPECIFICITY OF Philippia TO CORAL HOSTS a mainly dead skeleton resting on the sand. o Both the gastropod and the coral were kept Extrapolating from the Hawaiian habitat alive in an aquarium. The Philippia remained data, from the single feeding observation, from alive for about two months and was secretive, the fact that the proboscis is adapted for feed­ retracting into its shell whenever ordinary light ing on coelenterates, and from the much more was shone on it. Infrared light was found not thorough knowledge of the mode of life of to disturb the animal. Late one evening, under H eliacus, it seems probable that Philippia is infrared, the Philippia was seen with its pro- host-specific to stony corals. The world distri- FIG. 5. Incompletely developed veliger larva of Pbilip pia (Psi/axis) erebsi) (March) with the 4-lobed velum full y exten ded, show ing also the operculum and non-tentaculate eyes.
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